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WORKING CLASSIFICATION OF THE GASTROPODA 281
The phylogeny and classification of the Pulmonata is
controversial (Tillier, 1984; Starobogatov, 1989; Haszprunar&
Huber, 1990; Nordsieck, 1993a; Salvini-Plawen &
Steiner, 1995; Barker, 2001; Dayrat et al., 2001; Dayrat
& Tillier, 2002), In most analyses the monophyly of the
limnetic Hygrophila ("higher" Basommatophora) and the
terrestrial Stylommatophora is confirmed, whereas the
relationships of these groups and the more basal pulmonales
(often united in a paraphyletic group "Archaeopulmonata")
remain unclear We use the division of the
Pulmonata into Basommatophora (sensu stricto) and
Eupulmonata as proposed by Haszprunar & Huber
(1990) and the inclusion of the Systellommatophora in
the Eupulmonata as proposed by H. Nordsieck (1993a)
and supported by Tillier et al (1995), Barker (2001),
Dayrat et al. (2001 ) and Dayrat & Tillier (2002). How/ev-
er, according to the molecular phylogenetic analysis of
Dutra-Clarke et al. (2001 ) and Grande et al. (2004) the
Basommatophora (sensu stricto) as well as the Eupulmonata
are polyphyletic.
224 The monophyly of the Hygrophila (Chilinoidea + Acrolox-
oidea + Lymnaeoidea + Planorboidea) is supported by
the cladistic analysis of Barker (2001 ) and Dayrat et al.
(2001 ), whereas neither the monophyly of the Thalassophila
(= Amphiboloidea + Siphonarioidea) nor the mono-
phyly of the Basommatophora is supported by these
analysis. According to the molecular phylogenetic anal-
ysis of Grande et al. (2004), at least the Siphonarioidea
have to be transferred to the Opisthobranchia (the Am-
phiboloidea and the Hygrophila were not considered in
that study). Classification based on Hubendick (1978)
and Nordsieck (1993a) For alternative views see Bark-
er (2001), Starobogatov (1976), Starobogatov & Prozo-
rova (1990), Swiderski (1990) and J. Walker (1988).
22^ Inclusion of Acroreiidae in the SPF Siphonarioidea is ten-
tative following Zilch (1959) Examination (Bouchet & Le
Renard, unpubl.) of a specimen of Acroreia baylei, from
brackish-water deposits from the Paris Basin, is incon-
clusive (no discernible protoconch; one low, raised ridge
running internally from apical region to shell margin).
^2^ Placement of Scalaxinae in Lymnaeidae is tentative
following Nordsieck (1986b)
22^ There are several differences between the results of a
molecular phylogenetic analysis by Morgan et al. (2002)
and Hubendick's (1978) classification, which is used here
with nomenclatural adjustments. In the tree of Morgan
et al. (2002), Laevapex, Ferrissia and Amerianna form a
clade which is the sister group of the Planorbinae. Thus,
the Miratestini might be considered a distinct subfamily
and the name Laevapicinae and probably also "Gundlachiinae"
would be synonyms of Miratestinae
(instead of Bulinini). Morgan et al. (2002) called the clade
including Laevapex and Ferrissia Ancylidae, but Ancy-
lus (the only genus of the Ancylini sensu Hubendick),
which was included in the Planorbinae by Hubendick
(1978), was not examined by them. TheCoretini should
be transferred from the Bulininae sensu Hubendick to
the Planorbinae and might form a clade together with
the Biomphalariini and the Planorbulini The Coretini are
not monophyletic. The Drepanotrematinae are not re-
lated to the Biomphalariini, but form the sister clade of
all other Planorbinae, and, thus, should be considered
an indépendant tribe of the Planorbinae. Some of the
family-group taxa distinguished by Hubendick (1978)
were not examined by Morgan et al. (2002).
228 The "astonishing Odyssey" of Orygoceras, based on an
uncoiled type species from the Miocene of Croatia and
by different authors classified in the families Caecidae,
Hydrobiidae, Valvatidae, and Planorbidae, has been
summarized by Harzhauser et al. (2003). They classi-
fied the genus in the family Planorbidae, based on the
similarity of protoconchs of Orygoceras fuchsi (KittI,
1886), from the Miocene of central Europe, and of Miocene
species of Gyraulus.
229 Name based on wrongly identified genus.
230 Classification based on D. W. Taylor (2003).
231 Clade Eupulmonata - Trimusculoidea + Ellobioidea +
Otinoidea + Systellommatophora + Stylommatophora.
The cladistic analyses of Barker (2001) and Dayrat &
Tillier (2002) support a monophyletic group (Geophila)
including the Onchidioidea, Veronicelloidea and
Stylommatophora, whereas H. Nordsieck (1993a) con-
sidered the Ellobioidea the sister-group of the Stylommatophora.
Conversely, the molecular phylogenetic
analysis of Dutra-Clarke et al. (2001 ) and Grande et al.
(2004) has Eupulmonata polyphyletic (see also Note 223)
232 The Otinoidea were included in the Systellommatophora
by Haszprunar & Huber (1990) and Nordsieck (1993a).
However, according to the cladistic analyses of Barker
(2001 ) and Dayrat & Tillier (2002) they are not related
to the Systellommatophora (= Onchidioidea + Veronicelloidea)
and do not even belong to the Eupulmonata.
233 The Smeagolidae are related to the Otinidae accord-
ing to Tillier (1984), Tillier & Ponder (1992) and Barker
(2001), whereas they were classified as Onchidioidea
by Haszprunar & Huber (1990) and Nordsieck (1993a).
23* Classification of Ellobiidae after Prias Martins (1996).
235 The Zaptychiinae were classified as a subfamily of the
Ellobiidae by Wenz (1938) and Zilch (1959), and con-
sidered as a family of the Carychioidea by Starobogatov
(1976).
236 Monophyly supported by the analyses of Nordsieck
(1993a) and Barker (2001).
23*" The subfamilies distinguished by Hoffmann (1925) have
been rejected by Forcart (1953).
238 The phylogeny and classification of the Stylommatophora
is controversial (Solem, 1978; Schileyko, 1979a, 1998-
2003; Boss, 1982; Nordsieck, 1986b; Tillier, 1989;
Emberton, 1 991 b; Wade, Mordan & Clarke, 2001 Barker,
;
2001 ). For a comparison of different classifications see
Emberton et al. (1990). We consider the arguments of
Barker (2001 ) for a basal position of the Elasmognatha
in the Stylommatophora convincing.
239 = Heterurethra sensu lato - Succineoidea + Athoracoph-
oroidea. The monophyly of the Elasmognatha is also
supported by the molecular phylogenetic analysis of
Tillier et al. (1995), Wade, Mordan & Clarke (2001 ) and
Dutra-Clarke et al. (2001).